Botanica Marina Vol. 45, 2002, pp.


© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia,

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

E. Rintels (SUVA-A 3891. it was decided to revise records in the light of recent findings. a = axial cell.. Fig. P. C. Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. 1998. 1989. P. N’Yeurt (UPF 584). Material and Methods Specimens were collected by reef-walking.232 A. 2 Jul. SUVA-A 5438L). A. Faletose (SUVA-A 3894. SUVA-A 5436L). A. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. 3897). 19 Mar. – French Polynesia: Moorea. Figs 1–3. 1984. Taharaa.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. 5480L (cystocarpic). 2. 1998. Skelton. 3895. – Taiwan: Anon. Fig. leg.‘Al’ and ‘CG’ refer to specimens housed at IRD. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. Prepared slides were examined using a Zeiss compound microscope. Melanamansia glomerata. A. Okidomari. leg. leg. SUVA-A 1566 (isotype). 13 May 1982. leg. com. Punaauia. – Samoa: Palolo Deep Reserve. Most recently. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. Nouméa. Fig. A. 1997. 6 Jun. Amansia rhodantha. Payri (8/84. Palolo Deep. M. 3896. Amansia rhodantha (Beqa. otherwise as in Holmgren et al. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). 3893. P. Tanegashima Island. Photomicrographs were obtained using an Olympus BX-50 compound microscope. Rarotonga. 1. Enomoto (SAP). D. Kagoshima: 9 August 1959. Sections were made using a freezing microtome. N’Yeurt (UPF 347). N’Yeurt (SUVA-A 5128). leg. leg. A. 1999. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. leg. Tahiti. 1–5 = order of formation of pericentral cells. Afiti and D. A. 1998. . Sakai and S. Tahiti. M. leg. in EPHE). 1998. New Caledonia. Skelton and Dave [SUVA-A 1565 (holotype). snorkelling or SCUBA diving. 3892. 1998. Baba (SAP 056163). Y. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. P. SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). R. 3. 3898. showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. 9 Feb. Amansia rhodantha and Amansia glomerata. – Cook Islands: Ngatangiia. 3899). A. 18 Sep. (1981). a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. The results of these investigations are described below. developed and printed in the laboratory. Fiji. Baba (SAP 056160). 25 October 1984 (SAP 046978). 5481L (spermatangial)]. pers. Tombleson and T. Fiji. ps = pseudo-pericentral cells). leg. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. Skelton and Dave (SUVA-A 1567). 2 Jul. 11 Jul. U. M. Furusato. Okinoerabujima Island (60 km north of Okinawa). and a camera-lucida attachment was used to make anatomical drawings. leg. Skelton. leg. deep algal rim.Agardh to co-exist in the flora.

Table I presents the average results of morphological measurements carried out on Amansia rhodantha.). W. N’Yeurt (SUVA-A 346.Canipo Is.. leg. 16 Oct. E. – Philippines: Actin. A. 1965. W. anon. 1838–42. S. J. South (SUVA-A 5544. H. J. E. R. 1995. Cribb (SAP 051714). – Fiji: ‘Feejee’. leg. 1983. – Australia: Heron Island QLD. leg. 1995. 25 September 1999. leg. Masuda et al. Garrigue (CG 77c. 1069).. n.. A. 16 Nov.W. P. Oahu.d..leg. Yasawa Group. 1995. S. – Lord Howe Island: 18 Aug. 1876. Viti Levu. D.9 18. Hildebrandt (L 0108600).7 54. 1996: 472. fide G. SIPHILEXP-78 (US 088532). G. leg. leg. leg. Kasahara (SUVA-A 1012. 17 Jul. Agardh 1841: 26. Agardh). 1819.Negros Oriental. 15 Oct. Isaac (L 0108594). – Fiji: ‘Feejee’. 28 Nov. 10 July 1992. Wilkes Exploring Expedition (US 04057b) (doubtful record. SIPHILEXP-78 (US 088530). Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. leg. n.Calindagan Reef. Candelaria Zambales. Rukuruku Village.d. A. 25 Feb.2 53. 1954. Amansia rhodantha and Melanamansia glomerata surveyed in this study. leg. Richer de Forges (AL 288d-CG 77a). 10 May 1978. 1982. Amansia rhodantha: n. Negros Oriental. Beqa Lagoon. Average values of selected characters for Amansia paloloensis. Garrigue (CH1). B. A. 26 Nov. leg.d. R. Villeneuve (SUVA-A 1737). R127). leg. L 0108598).Siaton. Sumbawa. Figs 4–10 . leg. Isaac (L 0108595). 24 April 1996. A. 1979. 347).1 22. leg. 1973. Garrigue (SUVAA 5436L). Taxonomic results Amansia rhodantha (Harvey) J. 1979. leg. 84 (L 0108603).7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. Cordero and Tuangpalan (US 091395). Huisman (MURU. M. Kasahara (SUVA-A 1013. Suva Reef. Daida (US 086755). leg. 1838–42. H. Wilkes Exploring Expedition (US 04057a). c. leg. P. M de Robillard (L 0108599). G. N’Yeurt (SUVA-A 5437L). 3 Jun 1978. Oahu. Lectotype. Ningaloo. Nanuyalevu. Huisman (CWC 1997. Jul. Magnesia Virae.Cuyo. leg. leg. Villeneuve (SUVA-A 1774). J. Eagles Nest. A.Ilocos Norte. leg. Montebello Islands. W. C. 07 Oct. Unia. 3 and 5 March 1992. 1982. 9 Sept.. n. 84. 16 March 1991. 84. JH240). J. leg. Cannon (SAP 052062). AL 288b).Solong-On. J. figs 1–11 (as Amansia glomerata C. P. 1988. Chesterfield. leg. G. leg.d. Hapmak. B. J. – New Caledonia: Lajeu Sudouest. Potipot Is. leg. Huisman (MURU. – Kenya: Mombassa. Magruder (US 086459). – New Caledonia: Lajeu Sudouest. Dumaguete. Jul. South in lit.Palawan. R. 3 and 15 Oct. 17 Mar. Plongeurs IRD (AL 288a. 30 Jun. E. leg. 1980. 1985. Suva Barrier Reef. C. n. leg. leg. leg. 21/22-Sept. leg. Oahu. Carlson (SUVA-A 281). (MURU. Naukathura Island. M. Huisman (MURU. Tangangge. Silva et al. G. leg. Viti Levu. Yasawa Islands. leg.Negros Oriental. Ballou (SUVA-A 5438L).Moreland (US 50851). Norris 1988b: 211. Bahati. 1980. SIPHILEXP-78 (US 013760). SIPHILEXP-79 (US 088533). paloloensis and Melanamansia glomerata specimens examined in this study. Ti Tree. Barrow Island.11 Feb.Pagodpud.3 2. leg. leg. Negros Oriental. Jurien Bay WA.Revision of some Amansia and Melanamansia species 233 Table I.5 3. leg. (US 02890). Ovalau. B. Lopta. leg. Lawai Kai.6 3. 1984. Kauai. Kauai. see text). 1834). Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. Catanduanes.. Gaudichaud (‘Amansia glomerata’. Mauritius. SIPHILEXP-78 (US 088531). 3 Feb.leg.9 11. leg. Bay of Sanggar. Hanauma Bay Beach Park. 1997. 29 May 1978. CG 77b). Snellius-II (L 0108601). Farlow (L 0108597). leg. Hildebrandt (L 0108596. leg.d. R. J. figs 37–42.P. M. leg. 1067). 1876. 13 July 1972. 21 Sep.Siquijor Is. 18 March 1992.Dumaguete City. Cordero and Tuangpalan (US 091394). leg. Vatualailai. – Kenya: Mombassa. – Mozambique: Ponta Abril. leg. 2000: 188. 24 Nov. South (SUVA-A 2460). – Philippines:Gao-oa. Tantabiddi. Viti Levu. H. 1981. leg. W. C. – Indonesia: Cape Batu Kerapo.14 May 1979. Basay. Kipukai. R.23 May 1978. A. LD 42600 / 0772). 1985. 06 Sept.. Banilad. leg. in TCD (Telfair. Hakula.Malo. 14 March 1998. 1984. 6 Jun 1978. Skelton and G. Diani Beach. South (SUVA-A 5431). B262). 14 Feb. leg. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. Huisman. Barrow Island. 27 Dec. C.. M. 5 June 1967. Keats. Inhaca Peninsula. 10 Aug. Namada. NR 248). leg. leg. A. Snellius-II (L 0108602). B79). 1994 (SUVA-A 797). SIPHILEXP-78 (US 088534). Littler (US 163490). leg. Salayer.

showing axial cell (a) surrounded by five pericentral cells.234 A. showing both lax and dense growth forms. showing marginal serrations. Type material of Delesseria rhodantha Harvey. Secondary branchlet. 9. D. 8. 6. in TCD (scale bar = 2. Figs 4. Fig. US 02890 (scale bars = 3 mm). Surface view of segment cells of blade alae. N’Yeurt Figs 4–10. South Pacific (scale bar = 10 µm). in TCD (scale bar = 200 µm). . Habit of mature plant from the type locality. SUVA-A 5437L. Mauritius.5 mm). and a pair of budding branchlets from the midrib in the lower half. Fig. Fig. 7. R. 5. Rehydrated type material of Delesseria rhodantha Harvey. Fig. Indian Ocean. Type material of Delesseria rhodantha Harvey. Detail of tetrasporangial stichidia on lateral endogenous branch. Axial structure of blade. Amansia rhodantha. in TCD (scale bar = 25 µm). 10. from Rotuma Island.

SUVA-A 3892 (scale bar = 2. 14. Fig. 11. showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Compare with Fig. Detail of blades. . SUVA-A 1565 (scale bar = 25 µm). Fiji.Revision of some Amansia and Melanamansia species 235 Figs 11–16. Habit of mature plant. Holotype of Amansia paloloensis. showing lateral endogenous branches and midrib. Note marginal cystocarps (arrowheads).5 mm). Segment cells of blade alae in surface view. Detail of the holotype (SUVA-A 1565). SUVA-A 5479L (scale bar = 1 mm). Fig. SUVA-A 3892 (scale bar = 12 mm). 12. SUVA-A 5436L (scale bar = 50 µm). 15. Tetrasporangial stichidia of Amansia rhodantha from Beqa. 4. Fig. Detail of morphologically distinct mature rosette and corticated axis. Fig. 16. 13. with numerous rosette axes. Fig. Fig. Amansia paloloensis from Samoa.

118C–J. with a range of 3–6. D. and the apex is characteristically in-rolled. Abbott 1999: 404. pl.8–14. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. pl. 24. showing smooth margins and elongate shape. Lewis and Norris 1987: 22. Plant from Unia. fig. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. 19.4. AL288b (scale bar = 5 mm). Amansia paloloensis South et Skelton 1999: 247. 20. Kenya. Known distribution: Hawaiian Islands. Okamura 1900: 71. The stem of plants is thick and cartilaginous. 4B. Philippines. Norris 1995: 67. 436B. 20. Weber-van Bosse 1923: 369. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. they are up to 4 cells long and non-vesiculate. up to 100 (on average 40) mm high. New Caledonia. N’Yeurt 1996: 428. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. with a range of 1–5. 23.5 µm in diameter that lies adjacent to the axial cell (Figs 3. L 0108598 (scale bar = 700 µm). Mauritius. L 0108598 (scale bar = 5 mm). The main axis is irregularly branched. Fig. Figs 17–24 Basionym: Amansia glomerata C. Agardh) R. Agardh) R. containing up to 10 Figs 17–24. Tropical Pacific Ocean. 185. 21. e–i. Axial structure of plant from Lajeu. New Caledonia. 0. figs 2–14 (type locality: Palolo Deep Marine Reserve. 100–112 µm in diameter. Payri and Meinesz 1985: 511. Falkenberg 1901: 416. figs 1–13. Norris 1988b: 211. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). Blade margins are smooth. up to 35 mm long and 6 mm broad. 9). The central axial cell is 28–30 µm in diameter. surrounded by five pericentral cells. Tetrasporangial stichidia of plant from Oahu. Norris 1995: 67. fig. fig. fig. fig. stem-like and rigid in basal portions. N’Yeurt et al. Tetrasporangia occur in flattened. Figs 11–16 Misapplied names: Amansia glomerata C. to 0. Vietnam. 1. Fig. SUVA-A 5431 (scale bar = 50 µm). becoming narrower and disappearing towards the apex. 128. Silva et al. Kylin 1956: 544. Kenya. denuded below. 22. figs 14–29. fig. R. Samoa). East Africa. Tsuda and Wray 1977: 103. Apia. Note that the stichidia is twice laterally branched. Melanamansia glomerata. Individual tetrasporangia are spherical and tetrahedrally divided. – Melanamansia glomerata (C. Deciduous.8 mm in diameter. 212a. Detail of blade of plant from Mombassa. Ultimate branchlets are ecorticate. Blade of plant from Lajeu. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. Agardh 1824: 194. figs 127. figs 19–21.5–0. Norris 1995: 66) is usually 4. Isaac 1956: 188. 267. Fig. pl. in up to 10 regularly disposed pairs. Fig. South-East Asia. 18. Carposporangia are lanceolate. holotype in TCD. Coppejans and Millar 2000: 333 (unverified record). Womersley and Bailey 1970: 336.236 A. 6. Internal structure is cellular. Millar 1999: 523 (unverified record). Japan. pl. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. Yamada and Tanaka 1938: 86. XXV. 198. 25–42 × 83–125 µm. Trichoblasts are sparse. Agardh 1824: 247. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. figs 1–11. Fritsch 1945: 570. 7 mm broad. up to 750 µm long and 250 µm broad. Kenya. 1995: 150. A central midrib is present. Payri and N’Yeurt 1997: 897. Yoshida et al. 31 fig. composed of two overlapping layers of elongate hexagonal cells in transverse rows. New Caledonia. Garrigue and Tsuda 1988: 63. Cribb 1983: 106. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. Jaasund 1976: 131. . Fig. Melanamansia glomerata (C. 19). Secondary blades develop as successive pairs on the dorsal side of the midrib. Plant from Potipot Island. CG77a (scale bar = 1 mm). The thallus is rose-red. pl. Distribution: Tropical Indian Ocean. Fig. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. Plants are brownish-red. Blades of plant assumed to be from the Fiji Islands (see text). 6E (unverified record). and are initially vesiculate. E. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). Philippines. CG77a (scale bar = 10 µm). Plant from Mombassa. US 4057b (scale bar = 1 mm). with in-rolled leaf apex. plate CXXVI. 1996: 87. figs 1–3 (type locality: Cap Malheureux. 2. on average 53 mm high. US 091395 (scale bar = 5 mm). Millar et al. Isaac and Isaac 1968: 24. Marginal teeth are up to 1 mm long and 445 mm broad. 85–110 µm thick. New Caledonia. Fiji. 1987: 61. or with regular serrations (endogenous branches). 1999: 573. Dublin). and are crisp and lanceolate. Fig. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. Norris 1995: 66) is on average 4. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. 197. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. Fig. 1. each cell is about 20–20 × 87–100 µm in surface view. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. 17. 77–110 µm thick. E. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide.

In all cases. the colour of the Melanamansia plants was noticeably brownish-red. and stained the herbarium mounting paper. Spermatangial heads are curved and arranged in marginal clusters. . procarps and cystocarps occur on endogenous marginal branches.Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter.

and a cross section of the blade clearly shows the five pericentral cells around the axial ring. which is welldocumented and illustrated in the protologue of the species. The present author has re-examined the holotype. The reproductive and trichoblast anatomy of A. New Caledonia. While there is no doubt as to the generic identity of the latter specimen. and absence of pseudopericentral cells. In the Fiji Islands. 23) but the plant has not been collected in Fiji waters since. Vidalia. paloloensis showed no major difference in the range of characteristics among them. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. rhodantha from the type collection and localities in Mauritius. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. Smith (1979: 38–40) points out that the accounting and . N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. a re-examination of the lectotype of Amansia glomerata C. one of which (US 04057a) is Amansia rhodantha. M. D.). thicker. as well as Amansia species from Taiwan. the absence of blades organized in morphologically distinct tight axes around the parent axis. The habit and description of the holotype by Harvey (1834: 151. From this study. and these authors considered the lack of highly corticated axes. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. Masuda pers. fig. rhodantha from the Indian Ocean and other localities. and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11. paloloensis from A. 14) and corticated rosette axes (Fig. Characters such as the degree of rosette formation. rhodantha (South and Skelton 1999: 249). length of the blades.238 A. having tight rosettes as well as very lax. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. 12).S. Fig. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. The Wilkes collection consists of two plants. when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. Now that the Samoan species has been compared with genuine herbarium material of A. rhodantha as described and illustrated by Norris (1988b: 211). figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. with for instance A. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. R. The drawing of the holotype (South and Skelton 1999: 246. Fiji. Exploring Expedition. Rotuma. 3) which distinguishes that plant from A. it appears to be conspecific with the latter species and thus extends the status of A. between 1838 and 1842 (US 04057b. MURU 248). and Ceramium. indicating that the morphology of the species is influenced perhaps by environmental factors. rhodantha. Kenya. and a weakly developed midrib as characters distinguishing A. Kuetzingia. regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. paloloensis from the type locality. an additional field test to distinguish Amansia spp. while the other (US 04057b) is Melanamansia glomerata. rhodantha as a pan-tropical entity. apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). more coriaceous branchlets with less apparent midribs (this study). Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. plate CXXVI. 2) indeed shows a plant with loriform. generally smaller. More importantly. isotype and other collections of A. a comparison of material of A. paloloensis. Eastern Australia. Mozambique and Western Australia. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. the Cook Islands and French Polynesia (Table I) with A. the Philippines. while Melanamansia becomes dark brown. is comparable to A. However. and trichoblasts lacking protective vesicles when young (Norris 1988b).13). since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. These plants also had blades with well-developed midribs in the lower half (Fig. On the basis of these observations. rhodantha from Western Australia (MURU B262. loriform blades reminiscent of some specimens of A. paloloensis. Japan. Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. com. On the other hand. Amansia tends to bleach to translucent white. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U.

type of endogenous branching. However. When pseudo-pericentral cells were present. Payri and N’Yeurt 1997) were reexamined.). in New Caledonia and Kenya). which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. in the case of Amansia paloloensis). and hence the precise locality and date of this Melanamansia collection from Fiji is unknown. L 0108603). except the Hawaiian samples. calm waters (such as blue holes. In most instances. Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. when present). paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. some specimens were virtually indistinguishable from specimens of A. M. and is backed by other morphological and biochemical differences (Norris 1988b. rhodantha. glomerata occurs in the same habitat as Amansia rhodantha (e. environmental factors seem to play a role in determining the degree of rosette formation. The author has unsuccessfully tried to obtain material from the latter locality. Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. Also. fig. 2). Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined.e. they were clearly apparent and about half the size of pericentral cells (Fig. 1995). From an examination of the specimens at hand. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig. thus placing them in Amansia. while in Malaysia Masuda et al. 1) shows both the presence of tight and lax rosettes of secondary branches. which was the main reason for separating the species from A. 13). 1996. paloloensis. the description of the species states that branches have the same shape as the main axis (i. 19). the Philippines. irregularly dichotomous proliferations lack alar cells. Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. fig. an examination of the holotype and liquid-preserved and pressed isotypes of A. pl. Indian Ocean Amansia rhodantha also lacked such cells. 5. Norris (1988b: 211) reported that rosette axes in . and a main axis with alar cells not very distinct from secondary branches. Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. It would seem that perhaps shallow. both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. fig. pers. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). 7). and in some instances lacking altogether. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. both on the same plant and within individuals in similar populations. in the case of A. These ventral. older plants tend to have thicker. adding to the confusion between the two genera. M. in some cases. However. Masuda. perhaps an adaptation to the exposed habitats where the plants came from. it was observed that rosettes of blades along the secondary axes. so it cannot be confirmed at this stage if M. while younger plants have less cortication and more lax or absent rosettes of blades. CXXVI. more corticated secondary and primary axes with tight rosettes. which also had a consistently thicker thallus. All Amansia rhodantha specimens examined had fairly prominent. Wilson and Kraft 2000). and would seem to function as attachment haptera. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. and some samples from Kenya. Ecological studies would be needed to verify these hypotheses. the presence or absence of bladelets in rosettes. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a.g. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. Indeed. com. rhodantha. pseudo-pericentral cells were universally absent. and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. Fiji and New Caledonia (Table I). degree of marginal serrations and thallus flattening. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. 41) similarly report the absence of pseudo-pericentral cells in A.g.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. and found to lack pseudo-pericentral cells. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. Generally. (2000: 188. rhodantha from Mauritius and Western Australia (compare Figs 4. For instance. glomerata occurs in the Solomon Islands.

which was also recorded by Norris (1988b) for Hawaiian material. There appears to be no notable difference in the range of thallus size. only being more or less surrounded by cortication. com. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. rhodantha and A. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A.). and found them relatively constant. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). An examination of these elegant cells in species of Amansia (Figs 10. paloloensis specimens studied. R. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. which are in fact determinate branches covered by the pericentral cell derivatives. with the length of segments ranging from 80 to 120 µm. as well as between A. These differences reinforce the conspecificity of the Amansia rhodantha and A. rhodantha specimens observed in this study. it was noted that in Melanamansia the range was more restricted.3. paloloensis. . In A. 15). In Melanamansia it averages 4. blade dimensions.4 (Norris 1995). but the range is reportedly less than in Amansia rhodantha. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. with most specimens having segment lengths of about 100 µm. Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. Melanamansia glomerata from all localities studied have consistently smoother. in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. rhodantha studied. 25. 20). D. Cover cells usually totally cover the developing sporangia. However. From Table I.240 A. Fig. a number of preliminary conclusions can be made: A. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. 16) and Melanamansia show no particular difference. This also varied according to the degree of rosette formation and age of the plant. which was not seen in Amansia rhodantha (Figs 6. as well as the generic distinctness of Melanamansia glomerata. but can also occur apically. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows. as well as more corticated axes. B. This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. the number ranged from 3 to 7. usually 3 or 4 with an average of 3. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. showing the currently known distribution. as the axial ring was still discernible in all sections made. smaller and narrower blades.

Bot. Isaac.. E. Taxonomic notes on marine algae from Malaysia. Foreword. cxxvi. Lund. Schofield. Professor Willem Prud’homme van Reine (L – Rijksherbarium. 2000. W. M. 336 figs. Kylin.. and possibly other localities which are not yet adequately investigated. Masuda. Monographie 26. Nouv. (Hooker) 1: 147–157. Millar for his generous assistance in obtaining copies of rare publications. H. E.Vietnam and Malaysia (M. Myxophyceae. and A. 1824. 1987. Lord Howe Island (G. V. Norris. initially vesiculate and composed of up to 8 cells: Amansia rhodantha. K. South pers. 1834. J. Lunda (Lund). Mar. 1988. Dr R. and each genus is consistently differentiated from the other by a number of anatomical and biochemical characters. Phaeophyceae. Professors Michio Masuda and Tadao Yoshida (SAP – Hokkaido University. S. Bot. Jaasund. N. and J. Kato. . Dr John Parnell (TCD – Trinity College. New Caledonia and the Philippines (this study). Hommersand. 1981. S. Holmgren. trichoblasts non-vesiculate. 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(2000) confirm that the genus Amansia does exist in the Pacific Ocean. Marine Algae of the Southern Great Barrier Reef. Bot. Shimada. Phang. I. I. E. 35: 165–366. Fritsch. Tsuda. Isaac. Cribb. 1976. 27: 7–28. First edition. W. Bull. Skelton (SUVA – The University of the South Pacific). xiv + 939 pp. sparse. 7th edition. Coppejans. com. xvi + 754 pp. R.A. flora and vegetation. Accepted 17 January 2002. J. 43: 315–346. Cambridge University Press. M. Fauna und Flora des Golfes von Neapel.. Part 1. Bot. ser. Garrigue. K. Marine Red Algae of the Hawaiian Islands. E. J. Seven species of Rhodophyceae. com. 1901. 2 folded maps. Mar. E. Sci. M. 1963. C. General account of the environment. 17: 297–380.. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. C. Soc. 443–457. 24 pls. Dublin). Leiden). Sapporo). Berlin. with descriptions of some new and little known species. East Africa Nat. and R. 1841. A. II. in the Mauritius. pls v–vii.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. Masuda pers.. M. J. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. K. Nat. Kenya (this study). xv + 673 pp. xxxviii + 312 pp. Hist. Lund). 1809. G. Sci. Ms Susanna Riebe (LD – Botanical Museum. from “Cap Malheureux”. Die Gattungen der Rhodophyceen. Professor G. The Herbaria of the World.A. Phil. In historiam algarum symbolae. The Structure and Reproduction of the Algae. F. Figure 25 presents a world distribution of these two genera as it is currently known. Dr Claude Payri (UPF – Tahiti). E. Observations sur la physiologie des algues marines. 3. J. Isaac. Part I. Honolulu. pig- References Abbott. Index Herbariorum. Smiths. composed of up to 4 cells: Melanamansia glomerata. Lewis. Mar. C. 173 + (2) pp. 1945. xvi + 477 + [1] pp. 1842. Essais sur une classification des algues et des polypiers calcifères de Lamouroux. Falkenberg. iii + 159 pp. J. Norris is thanked for his helpful insights and suggestions for this research. Handbook No 2. pls cxxv. ments rose-red. Sci. 1999. Notice of a collection of algae. Gleerups Förlag. 29: 1–38. 22: 161–193. P. J. Australian Coral Reef Society: Brisbane. Intertidal Seaweeds in Tanzania. and F. trichoblasts on every second or third axial cell. Millar. Marine red algae from the North coast of Papua New Guinea. Marine botany of the Kenya coast. Rhodophyceae. Bohn. Agardh. Scheltema & Holkema. Utrecht. Marine algae of Inhaca Island and of the Inhaca Peninsula. S.. Volume II. K. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). Cal. Harvey. Agardh. Contrib. H. K. J. Paris 1: 330–333. 10 figs. 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